{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,30]],"date-time":"2026-04-30T02:56:17Z","timestamp":1777517777262,"version":"3.51.4"},"reference-count":69,"publisher":"Wiley","license":[{"start":{"date-parts":[[2026,4,9]],"date-time":"2026-04-09T00:00:00Z","timestamp":1775692800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,4,9]],"date-time":"2026-04-09T00:00:00Z","timestamp":1775692800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100008567","name":"Leeds Beckett University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100008567","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["physoc.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Journal of Physiology"],"abstract":"<jats:sec>\n                    <jats:title>Abstract<\/jats:title>\n                    <jats:p>\n                      The characterization of skeletal muscle phenotypes in diving populations remains one of the least explored domains of breath\u2010hold physiology, representing a critical gap in our understanding of how skeletal muscle adapts to the unique demands of breath\u2010hold diving. Accordingly the present study investigated specific markers of skeletal muscle structure and metabolism in competitive breath\u2010hold divers. Twenty males volunteered to participate in this study (10 competitive breath\u2010hold divers; 10 non\u2010divers), matched for age, body size and whole\u2010body aerobic capacity (). A percutaneous skeletal muscle biopsy was obtained from the\n                      <jats:italic>m. vastus lateralis<\/jats:italic>\n                      to quantify capillarization, fibre\u2010type distribution (i.e. types I, IIa and II other), protein content of mitochondrial complexes, monocarboxylate transporter (MCT) isoforms and citrate synthase activity. MCT4 content was 28% higher in breath\u2010hold divers compared to non\u2010divers (\n                      <jats:italic>P<\/jats:italic>\n                      \u00a0=\u00a00.020), whereas MCT1 and citrate synthase activity showed no between\u2010group differences (\n                      <jats:italic>P<\/jats:italic>\n                      \u2265 0.161). Complex V content was higher in the non\u2010divers (\n                      <jats:italic>P<\/jats:italic>\n                      \u00a0=\u00a00.049), whereas no between\u2010group differences were noted for complexes I, II, III and IV (\n                      <jats:italic>P<\/jats:italic>\n                      \u2265 0.253). Capillarization was significantly higher in breath\u2010hold divers (\n                      <jats:italic>P<\/jats:italic>\n                      \u2264 0.048), whereas fibre\u2010type distribution did not differ between groups (\n                      <jats:italic>P<\/jats:italic>\n                      \u00a0=\u00a00.999). Competitive breath\u2010hold divers exhibited skeletal muscle characteristics indicative of enhanced blood\u2013muscle exchange capacity and augmented lactate efflux potential. Such adaptations may confer an advantage during prolonged breath\u2010holds by preserving glycolytic function and maintaining redox homeostasis. In recovery these traits likely facilitate more efficient clearance of metabolic byproducts.\n                      <jats:boxed-text content-type=\"graphic\" position=\"anchor\">\n                        <jats:graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" mimetype=\"image\/png\" position=\"anchor\" specific-use=\"enlarged-web-image\" xlink:href=\"graphic\/tjp70497-gra-0001-m.png\">\n                          <jats:alt-text>image<\/jats:alt-text>\n                        <\/jats:graphic>\n                      <\/jats:boxed-text>\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Key points<\/jats:title>\n                    <jats:p>\n                      <jats:list list-type=\"bullet\">\n                        <jats:list-item>\n                          <jats:p>The skeletal muscle phenotype of breath\u2010hold divers remains poorly characterized, limiting understanding of how skeletal muscle adapts to the physiological demands of prolonged breath\u2010holding.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>This study compared skeletal muscle structure and metabolic markers between competitive breath\u2010hold divers and non\u2010diving controls matched for age, body size and whole\u2010body aerobic capacity.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>Breath\u2010hold divers exhibited greater skeletal muscle capillarization, indicating an enhanced capacity for blood\u2013muscle exchange.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>They also showed an increased potential to remove lactate from skeletal muscle tissue.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>These adaptations likely support sustained glycolytic function and redox balance during prolonged breath\u2010holds, while facilitating more efficient clearance of metabolic byproducts during recovery.<\/jats:p>\n                        <\/jats:list-item>\n                      <\/jats:list>\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1113\/jp290732","type":"journal-article","created":{"date-parts":[[2026,4,9]],"date-time":"2026-04-09T17:55:21Z","timestamp":1775757321000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Diving into the unknown: Evidence of enhanced skeletal muscle lactate efflux potential and blood\u2013muscle exchange in competitive breath\u2010hold divers"],"prefix":"10.1113","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5991-0733","authenticated-orcid":false,"given":"Antonis","family":"Elia","sequence":"first","affiliation":[{"name":"Division of Environmental Physiology, Department of Physiology and Pharmacology Karolinska Institute  Stockholm Sweden"},{"name":"Carnegie School of Sport Leeds Beckett University  Leeds UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1422-0154","authenticated-orcid":false,"given":"Matthew","family":"Lees","sequence":"additional","affiliation":[{"name":"Exercise Metabolism Research Group, Department of Kinesiology McMaster University  Hamilton Ontario Canada"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"John","family":"O'Hara","sequence":"additional","affiliation":[{"name":"Carnegie School of Sport Leeds Beckett University  Leeds UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Nathan","family":"Hodson","sequence":"additional","affiliation":[{"name":"Department of Sport and Exercise Science, Institute of Sport Manchester Metropolitan University  Manchester UK"},{"name":"School of Sport, Exercise and Rehabilitation Sciences University of Birmingham  Birmingham UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-9808-5519","authenticated-orcid":false,"given":"Jordan","family":"Acheson","sequence":"additional","affiliation":[{"name":"Department of Sport and Exercise Science, Institute of Sport Manchester Metropolitan University  Manchester UK"},{"name":"School of Sport, Exercise and Rehabilitation Sciences University of Birmingham  Birmingham UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Oliver","family":"Wilson","sequence":"additional","affiliation":[{"name":"Carnegie School of Sport Leeds Beckett University  Leeds UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-3463-2669","authenticated-orcid":false,"given":"Paul","family":"Parker","sequence":"additional","affiliation":[{"name":"Royal Centre for Defence Medicine  Birmingham UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7291-9516","authenticated-orcid":false,"given":"Matthew","family":"Barlow","sequence":"additional","affiliation":[{"name":"Carnegie School of Sport Leeds Beckett University  Leeds UK"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"311","published-online":{"date-parts":[[2026,4,9]]},"reference":[{"key":"e_1_2_6_2_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1985.sp015794"},{"key":"e_1_2_6_3_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature06613"},{"key":"e_1_2_6_4_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1365-201X.2003.01185.x"},{"key":"e_1_2_6_5_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajpendo.1998.274.1.E102"},{"key":"e_1_2_6_6_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajpendo.2000.279.5.E1131"},{"key":"e_1_2_6_7_1","doi-asserted-by":"publisher","DOI":"10.1152\/jappl.1996.81.1.355"},{"key":"e_1_2_6_8_1","doi-asserted-by":"publisher","DOI":"10.1152\/jappl.1966.21.1.33"},{"key":"e_1_2_6_9_1","doi-asserted-by":"publisher","DOI":"10.1093\/geroni\/igz036"},{"key":"e_1_2_6_10_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajpregu.00503.2006"},{"issue":"16","key":"e_1_2_6_11_1","first-page":"2947","article-title":"High fatty acid oxidation capacity and phosphorylation control despite elevated leak and reduced respiratory capacity in northern elephant seal muscle mitochondria","volume":"217","author":"Chicco A. J.","year":"2014","journal-title":"Journal of Experimental Biology"},{"key":"e_1_2_6_12_1","first-page":"37","article-title":"Regulation of pyruvate metabolism in mammalian tissues","volume":"15","author":"Denton R. M.","year":"1979","journal-title":"Essays in Biochemistry"},{"key":"e_1_2_6_13_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00374176"},{"key":"e_1_2_6_14_1","doi-asserted-by":"publisher","DOI":"10.1113\/EP088404"},{"key":"e_1_2_6_15_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajpregu.00260.2023"},{"key":"e_1_2_6_16_1","doi-asserted-by":"publisher","DOI":"10.1152\/japplphysiol.00508.2025"},{"key":"e_1_2_6_17_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00421-021-04664-x"},{"key":"e_1_2_6_18_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajpregu.00171.2022"},{"key":"e_1_2_6_19_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00421-025-05752-y"},{"key":"e_1_2_6_20_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00421-019-04230-6"},{"key":"e_1_2_6_21_1","doi-asserted-by":"publisher","DOI":"10.1152\/jappl.1997.82.2.447"},{"key":"e_1_2_6_22_1","doi-asserted-by":"publisher","DOI":"10.1016\/S1095-6433(03)00134-X"},{"key":"e_1_2_6_23_1","doi-asserted-by":"publisher","DOI":"10.1152\/jappl.1997.83.4.1282"},{"issue":"2","key":"e_1_2_6_24_1","first-page":"81","article-title":"Electrocardiogram during deep breath\u2010hold dives by elite divers","volume":"18","author":"Ferrigno M.","year":"1991","journal-title":"Undersea Biomedical Research"},{"key":"e_1_2_6_25_1","doi-asserted-by":"publisher","DOI":"10.1002\/j.2040-4603.1995.tb01723.x"},{"key":"e_1_2_6_26_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1600-0838.2005.00440.x"},{"key":"e_1_2_6_27_1","first-page":"19","volume-title":"Mitochondrial Physiology Network. 19.12","author":"Gnaiger E.","year":"2014"},{"key":"e_1_2_6_28_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF02691277"},{"key":"e_1_2_6_29_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajpheart.2000.279.6.H3146"},{"key":"e_1_2_6_30_1","doi-asserted-by":"publisher","DOI":"10.1152\/jappl.1968.25.5.542"},{"key":"e_1_2_6_31_1","doi-asserted-by":"publisher","DOI":"10.1139\/h97-002"},{"key":"e_1_2_6_32_1","doi-asserted-by":"publisher","DOI":"10.1007\/s004240050273"},{"key":"e_1_2_6_33_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1908.sp001258"},{"issue":"5","key":"e_1_2_6_34_1","first-page":"433","article-title":"Daily diving pattern of Korean and Japanese breath\u2010hold divers (ama)","volume":"18","author":"Hong S. K.","year":"1991","journal-title":"Undersea Biomedical Research"},{"key":"e_1_2_6_35_1","doi-asserted-by":"publisher","DOI":"10.1152\/jappl.1963.18.3.457"},{"key":"e_1_2_6_36_1","doi-asserted-by":"publisher","DOI":"10.1016\/0167-4889(94)00209-W"},{"key":"e_1_2_6_37_1","doi-asserted-by":"publisher","DOI":"10.1152\/physrev.1997.77.2.321"},{"key":"e_1_2_6_38_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1365-201X.1998.0305f.x"},{"key":"e_1_2_6_39_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7793.1999.0633s.x"},{"key":"e_1_2_6_40_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0201401"},{"key":"e_1_2_6_41_1","doi-asserted-by":"publisher","DOI":"10.14814\/phy2.12033"},{"key":"e_1_2_6_42_1","doi-asserted-by":"publisher","DOI":"10.1152\/japplphysiol.01474.2005"},{"key":"e_1_2_6_43_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1919.sp001844"},{"issue":"6","key":"e_1_2_6_44_1","first-page":"463","article-title":"Alveolar gas composition before and after maximal breath\u2010holds in competitive divers","volume":"33","author":"Lindholm P.","year":"2006","journal-title":"Undersea & Hyperbaric Medicine: Journal of the Undersea and Hyperbaric Medical Society, Inc"},{"key":"e_1_2_6_45_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajpregu.90564.2008"},{"key":"e_1_2_6_46_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0095092"},{"key":"e_1_2_6_47_1","doi-asserted-by":"publisher","DOI":"10.1002\/jcsm.12410"},{"key":"e_1_2_6_48_1","doi-asserted-by":"publisher","DOI":"10.1152\/japplphysiol.00624.2019"},{"issue":"12","key":"e_1_2_6_49_1","first-page":"1123","article-title":"Muscle characteristics in career breath\u2010hold divers: Effect of water temperature","volume":"76","author":"Park J. B.","year":"2005","journal-title":"Aviation Space and Environmental Medicine"},{"key":"e_1_2_6_50_1","doi-asserted-by":"publisher","DOI":"10.1139\/H08-097"},{"key":"e_1_2_6_51_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajpendo.1999.276.2.E255"},{"issue":"5","key":"e_1_2_6_52_1","first-page":"E843","article-title":"Distribution of the lactate\/H+ transporter isoforms MCT1 and MCT4 in human skeletal muscle","volume":"276","author":"Pilegaard H.","year":"1999","journal-title":"American Journal of Physiology"},{"key":"e_1_2_6_53_1","doi-asserted-by":"publisher","DOI":"10.17305\/bb.2024.10787"},{"key":"e_1_2_6_54_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.metabol.2004.12.007"},{"key":"e_1_2_6_55_1","doi-asserted-by":"publisher","DOI":"10.1152\/jappl.1993.75.4.1911"},{"key":"e_1_2_6_56_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-4615-1875-4_91"},{"key":"e_1_2_6_57_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajpheart.1999.277.6.H2247"},{"key":"e_1_2_6_58_1","doi-asserted-by":"publisher","DOI":"10.1016\/0002-9149(85)91054-9"},{"issue":"1","key":"e_1_2_6_59_1","first-page":"27","article-title":"Underwater working times in two groups of traditional apnea divers in Asia: The Ama and the Bajau","volume":"41","author":"Schagatay E.","year":"2011","journal-title":"Diving and Hyperbaric Medicine"},{"key":"e_1_2_6_60_1","doi-asserted-by":"publisher","DOI":"10.1038\/nmeth.2019"},{"key":"e_1_2_6_61_1","doi-asserted-by":"publisher","DOI":"10.1213\/ANE.0000000000002864"},{"key":"e_1_2_6_62_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0006-2952(99)00292-0"},{"issue":"1","key":"e_1_2_6_63_1","first-page":"37","article-title":"Is hypoxia\u2010inducible factor (HIF\u20101) alpha involved in taravana syndrome?","volume":"94","author":"Sponsiello N.","year":"2021","journal-title":"Journal of Biological Research"},{"issue":"5","key":"e_1_2_6_64_1","first-page":"361","article-title":"Breath\u2010hold duration in man and the diving response induced by face immersion","volume":"15","author":"Sterba J. A.","year":"1988","journal-title":"Undersea Biomedical Research"},{"key":"e_1_2_6_65_1","doi-asserted-by":"publisher","DOI":"10.1096\/fasebj.21.6.LB115"},{"key":"e_1_2_6_66_1","doi-asserted-by":"publisher","DOI":"10.1002\/mus.21945"},{"key":"e_1_2_6_67_1","doi-asserted-by":"publisher","DOI":"10.1152\/jappl.1990.68.6.2369"},{"key":"e_1_2_6_68_1","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M511397200"},{"key":"e_1_2_6_69_1","doi-asserted-by":"publisher","DOI":"10.1152\/jappl.2001.91.1.173"},{"key":"e_1_2_6_70_1","doi-asserted-by":"publisher","DOI":"10.1152\/jappl.1993.75.1.264"}],"container-title":["The Journal of Physiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/pdf\/10.1113\/JP290732","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/full-xml\/10.1113\/JP290732","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/pdf\/10.1113\/JP290732","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,12]],"date-time":"2026-04-12T01:43:54Z","timestamp":1775958234000},"score":1,"resource":{"primary":{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/10.1113\/JP290732"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,4,9]]},"references-count":69,"alternative-id":["10.1113\/JP290732"],"URL":"https:\/\/doi.org\/10.1113\/jp290732","archive":["Portico"],"relation":{},"ISSN":["0022-3751","1469-7793"],"issn-type":[{"value":"0022-3751","type":"print"},{"value":"1469-7793","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,4,9]]},"assertion":[{"value":"2025-12-30","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-03-10","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-04-09","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"JP290732"}}